Veget Hist Archaeobot (2014) 23:607–613
DOI 10.1007/s00334-013-0414-2
ORIGINAL ARTICLE
Forage quality of leaf-fodder from the main broad-leaved woody
species and its possible consequences for the Holocene
development of forest vegetation in Central Europe
Pavla Hejcmanová • Michaela Stejskalová •
Michal Hejcman
Received: 1 April 2013 / Accepted: 25 August 2013 / Published online: 5 September 2013
Ó Springer-Verlag Berlin Heidelberg 2013
Abstract Leaf-hay was the principal winter feed of
livestock from the Neolithic until the first archaeological
records of scythes dated to the Iron Age (700–0 B.C.).
Despite the use of meadow hay, leaf-fodder remained an
important winter supplement until the present. Archaeo-
logical evidence lists Quercus, Tilia, Ulmus, Acer, Fraxi-
nus and Corylus as woody species harvested for leaf-
fodder, while Fagus, Populus or Carpinus were rarely
used. The aim of our study was to test whether the use of
listed woody species followed the pattern of their forage
quality (syn. nutritive value). In late May 2012, we col-
lected leaf biomass at four localities in the Czech Republic
and determined concentrations of N, P, K, Ca, Mg, neutral-
and acid-detergent fibre and lignin. Species with leaves of
low forage quality were Carpinus betulus, Fagus sylvatica
and Quercus robur, species with leaves of intermediate
quality were Corylus avellana and Populus tremula and
species with leaves of high quality were Ulmus glabra,
Fraxinus excelsior, Tilia cordata and Acer platanoides.
Selective browsing and harvesting of high quality species
Acer, Fraxinus, Tilia and Ulmus thus probably supported
their decline in the Bronze and Iron ages and supported the
expansion of Carpinus and Fagus. Our results indicate that
Communicated by K.-E. Behre.
P. Hejcmanová (&)
Faculty of Tropical AgriSciences, Czech University of Life
Sciences, Kamýcká 129, Prague 6-Suchdol 16521, Czech
Republic
e-mail: hejcmanova@ftz.czu.cz
M. Stejskalová  M. Hejcman
Faculty of Environmental Sciences, Czech University of Life
Sciences, Kamýcká 1176, Prague 6-Suchdol 16521, Czech
Republic
our ancestors’ practice of exploiting woody species as leaf-
hay for winter fodder followed their nutritive value.
Keywords Agricultural history  Leaf-fodder 
Livestock feeding  Nutritive value  Prehistory
Introduction
The practice of collecting leaves and twigs for livestock
feeding is probably the oldest method of fodder harvesting
in Europe and is still widely practiced in tropical and
subtropical regions of Asia and Africa (e.g. Roothaert and
Paterson 1997; Le Houerou 2000; Cheema et al. 2011).
Leaf-hay (syn. leaf-fodder) played a significant role in
livestock feeding especially during the winter at least from
the Late Neolithic (3000 B.C.) and was widely used over the
course of time in Europe (Dreslerová 2012). The first
written records about the use of leaf-fodder come from
Roman authors, Marcus Porcius Cato (234–149 B.C., De
Agricultura, Hooper and Ash 1935) or Lucius Junius
Moderatus Columella (A.D. 4(?)-70 A.D., De Re Rustica,
Ash 1941). Another valuable source of knowledge of pre-
historic livestock foddering with twigs and leaves comes
mainly from charcoal (Regnell 2003), and from analyses of
plant macrofossils and pollen from sheep and goat copro-
lites found in the Neolithic lake-shore settlements in the
Alps (Akeret and Jacomet 1997; Akeret et al. 1999) or from
the Neolithic shepherd’s shelter of la Grande Rivoire
(Delhon et al. 2008; Martin 2011).
The advantage of leaf-hay is that it can be very effi-
ciently collected without the use of any special tools such
as sharp metal scythes or sickles. Based on recent analogies
(Austad 1988) it is supposed that there have been three
main techniques of leaf-hay collection since the Neolithic:
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(1) Pollarding, which consists in cutting off the top of the
tree and harvesting the shoots.
(2) Shredding, which leaves the trunk and crown intact
while the side branches are removed.
(3) Coppicing, based on cutting suckers or young shoots
directly from the tree base.
Leaf-hay was widely used across the whole of Europe,
from Greece (Halstead 1998) to Great Britain (Smith 2010)
and from Italy and France (Haas et al. 1998; Karg 1998) to
Scandinavia (Slotte 2001). In each region, different woody
species were used according to the regional vegetation and
woody species availability, which latter could be ensured
for instance by transhumance (Akeret and Jacomet 1997;
Poschlod and WallisDeVries 2002). Many archaeological
records have confirmed year-round leaf-hay foddering of
sheep and goats in stables in the Alps, using ash (Fraxinus
excelsior), lime (Tilia spp.) and hazel (Corylus avellana),
or fir (Abies alba) (Rasmussen 1989; Rasmussen 1993;
Delhon et al. 2008). In the northern part of Europe woody
species exploited as fodder for cattle, horses, sheep and
goats were ash, alder (Alnus glutinosa), birch (Betula spp.),
lime, poplar (Populus spp.) and several coniferous species,
primarily Scots pine (Pinus sylvestris) or juniper (Junipe-
rus communis) (Austad 1988; Austad and Hauge 2006).
However, the most important tree species for leaf-hay
across Europe, considered of a high forage quality, was the
elm (Ulmus spp.). Consequently the elm has been sug-
gested as having been selectively harvested by Neolithic
farmers. This could have resulted, according to pollen
analysis, in the marked elm decline (Troels-Smith 1960;
Iversen 1973; Magyari et al. 2012). On the other hand,
hornbeam (Carpinus betulus) or beech (Fagus sylvatica)
were not exploited to the same extent due to being con-
sidered of low forage quality (Halstead 1998).
Although it is supposed that different woody species
were selected for leaf harvesting according to their quality,
the forage qualities of common broad-leaved woody spe-
cies in Central Europe has never been compared with each
other. Forage quality can be evaluated according to con-
centration of macro-nutrients, N, P, K, Ca and Mg partic-
ularly, their ratios, Ca:P and K:(Ca?Mg) and by fibre
content (cellulose, hemi-cellulose and lignin) which pre-
determine organic matter digestibility (Hejcman et al.
2006, 2010; Pavlů et al. 2006).
In this paper, we investigated whether the main Euro-
pean broad-leaved woody species Acer platanoides, C.
betulus, C. avellana, Fagus sylavtica, F. excelsior, Quercus
robur, Populus tremula, Tilia cordata and Ulmus glabra
differ in their forage quality for livestock, and consequently
whether their supposed use could follow the pattern of this.
In addition, we discussed possible consequences of forage
quality of different woody species for their selection by
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Veget Hist Archaeobot (2014) 23:607–613
livestock and for the woody species composition of agri-
culturally used forests in Central Europe.
Materials and methods
Biomass sampling
For the analysis, we selected nine broad-leaved woody
species common in Central Europe at least since the
Bronze Age (1900–800 B.C.). We collected the leaf biomass
(fully expanded leaves, blades and petioles together) of
selected woody species from at least three individuals of
each species at each of four selected localities in late May
2012. These localities were broad-leaved forests and their
margins in Bohemia, the western part of the Czech
Republic:
(1) Kozly (50° 150 N, 14° 320 E, 167 m a.s.l.);
(2) Měšice (50° 110 N, 14° 310 E, 203 m a.s.l.);
(3) Běstvina (49°500 N, 15°350 E, 338 m a.s.l.);
(4) Mšec (50°120 N, 13°520 E, 435 m a.s.l.).
Mean annual precipitation and temperatures ranged at
the sampling sites from 614 to 723 mm and from 8 to 9 °C,
respectively. We collected in total 36 (nine species 9 four
replicates) leaf biomass samples, which were then oven-
dried at 60 °C for 48 h and ground to powder.
Leaf chemical properties
In the leaf samples, the concentrations of macro-elements
(N, P, K, Ca, Mg) and the residual ash content (ash–
(P?K?Ca?Mg)) neutral- (NDF) and acid-detergent fibre
(ADF) and acid-detergent lignin (ADL) were determined.
NDF represents cellulose, hemi-cellulose and lignin toge-
ther, ADF represents cellulose and lignin. The N concen-
tration in the plant samples was determined using an
automated analyser TruSpec (LECO Corporation, USA) by
combustion with oxygen in an oven at 950 °C. Combustion
products were mixed with oxygen and the mixture passed
through an infrared CO2 detector and through a circuit for
aliquot ratio where carbon is measured as CO2. Gases in the
aliquot circuit were transferred into helium as a carrying
gas, conducted through hot copper and converted to N.
Biomass samples were burnt in a microwave oven at
temperature of 550 °C and weighed in order to determine
ash content. Biomass samples were mineralized using aqua
regia and P, K, Ca and Mg concentrations were then
determined in the solution using ICP-OES (Varian Vist-
aPro, Mulgrave, Vic., Australia). Residual ash containing
mostly Si was calculated as the ash content minus the sum
of P, K, Ca and Mg concentrations. NDF, ADF and ADL
Veget Hist Archaeobot (2014) 23:607–613 609

contents were determined by standard methods of AOAC
(1984).
All analyses were performed in the accredited national
laboratory Ekolab Žamberk (http://www.ekolab.zamberk.
cz). N:P, Ca:P and K:(Ca?Mg) ratios were calculated from
determined concentrations.
Data analyses
The data were tested by the Kolmogorov–Smirnov test of
normality and met the assumption for the use of parametric
tests. One-way ANOVA followed by post hoc comparison
using the Tukey’s multiple range tests in the STATISTICA
9.0 program (StatSoft, Tulsa, USA) were used to identify
significant differences in concentrations of nutrients and
NDF, ADF and ADL contents among species.
Unconstrained principal component analysis (PCA) in
the CANOCO for Windows 4.5 program (Ter Braak and
Šmilauer 2002) was used to analyze relationships among
leaf chemical properties and similarity of 36 samples. Data
were log-transformed before the analysis. The results of the
PCA analysis were visualized in the form of an ordination
diagram constructed by the CanoDraw program (Ter Braak
and Šmilauer 2002).
Results
No significant effect of species on N and Mg concentra-
tions in leaves was recorded (P [ 0.05, Table 1). Nitrogen
concentration ranged from 28.2 to 36.0 g kg-1 and Mg
from 2.6 to 4.1 g kg-1, respectively. Effect of species on
concentrations of other elements in leaves was significant
(all P \ 0.01).
Concentration of P ranged from 1.97 g kg-1 in
Carpinus to 3.22 g kg-1 in Fraxinus, concentration of K
ranged from 9.1 g kg-1 in Fagus to 20.9 g kg-1 in Frax-
inus and concentration of Ca ranged from 9.3 g kg-1 in
Quercus to 17.2 g kg-1 in Fraxinus.
There was a significant effect of species on N:P, Ca:P
and K:(Ca?Mg) ratios (all P \ 0.001) in leaves. N:P ratio
ranged from 10.5 in Fraxinus to 15.5 in Carpinus, Ca:P
ratio ranged from 3.8 in Quercus to 7.2 in Corylus, and
K:(Ca?Mg) ratio ranged from 0.69 in Carpinus to 1.46 in
Tilia.
There was a significant effect of species on content of
NDF, ADF, ADL and residual ash in leaves (all P \ 0.001,
Table 2). Content of NDF ranged from 397 g kg-1 in Ul-
mus to 663 g kg-1 in Fagus, content of ADF ranged from
243 g kg-1 in Ulmus to 480 g kg-1 in Fagus, content of
ADL ranged from 85 g kg-1 in Carpinus to 236 g kg-1 in
Fagus and content of residual ash ranged from 24.8 g kg-1
in Quercus to 51.8 g kg-1 in Ulmus.
Results of the PCA analysis are presented in the form of
the ordination diagram in Fig. 1. The first ordination axis
explained 32 %, the first two axes together 56 % and the
first four axes together 84 % variability of leaf chemical
data. Concentrations of macro-elements were positively
correlated one with another and generally bore no relation
to the contents of NDF, ADF and ADL which were posi-
tively correlated with one another and negatively correlated
with the K:(Ca?Mg) ratio. The first axis divided species

Table 1 Concentration (mean ± standard error of mean) of N, P, K, Ca, Mg and N:P, Ca:P and K:(Mg?Ca) ratios in leaf-fodder of the studied
species
Species N (g kg-1) P (g kg-1) K (g kg-1) Ca (g kg-1) Mg(g kg-1) N:P ratio Ca:P ratio K:(Mg ? Ca)

Acer platanoides 28.2 ± 1.2a 2.7 ± 0.3a,b 18.0 ± 0.6a,b 12.3 ± 0.5a,b,c 3.0 ± 0.2a 10.8 ± 1.3a,b 4.6 ± 0.4a,b 1.18 ± 0.07a,b
a a c,d a,b,c a c a,b
Carpinus betulus 30.7 ± 0.6 1.9 ± 0.1 10.6 ± 1.0 12.7 ± 1.7 2.7 ± 0.4 15.5 ± 0.4 6.4 ± 0.7 0.69 ± 0.04a
a a,b a,b,d b,c a a,b,c b
Corylus avellana 31.5 ± 1.4 2.3 ± 0.1 16.0 ± 1.9 16.4 ± 1.3 3.6 ± 0.3 13.8 ± 1.2 7.2 ± 0.8 0.79 ± 0.06a
Fagus sylvatica 31.4 ± 2.9a 2.1 ± 0.2a 9.1 ± 0.5c 10.2 ± 0.4a,b 2.6 ± 0.2a 15.3 ± 0.3b,c 5.1 ± 0.3a,b 0.71 ± 0.05a
a b a c a a a,b
Fraxinus excelsior 33.2 ± 2.1 3.2 ± 0.2 20.9 ± 0.9 17.2 ± 1.3 4.1 ± 0.7 10.5 ± 1.0 5.4 ± 0.6 0.99 ± 0.05a,b
a a,b a,b a,b,c a a,b,c a,b
Populus tremula 28.9 ± 1.0 2.6 ± 0.2 18.3 ± 2.0 15.4 ± 1.1 2.8 ± 0.4 11.3 ± 0.7 6.1 ± 0.6 1.01 ± 0.09a,b
a a,b b,c,d a a a,b,c a
Quercus robur 32.9 ± 0.9 2.5 ± 0.1 11.6 ± 0.7 9.3 ± 0.5 2.6 ± 0.2 13.5 ± 0.8 3.8 ± 0.1 0.97 ± 0.03a,b
a a,b a a,b,c a a,b,c a
Tilia cordata 36.0 ± 1.2 3.0 ± 0.5 19.7 ± 0.4 12.1 ± 2.4 2.8 ± 0.6 12.6 ± 1.3 4.1 ± 0.7 1.46 ± 0.25b
Ulmus glabra 34.3 ± 3.3a 2.7 ± 0.1a,b 19.9 ± 2.7a 14.2 ± 1.9a,b,c 2.7 ± 0.3a 12.7 ± 0.7a,b,c 5.2 ± 0.5a,b 1.18 ± 0.08a,b
Meadow hay 20.0–28.7 2.7–3.7 24.1–34.0 6.0–8.4 1.5–4 5–10 1.7–2.5 –
Optimum range 19.2–25.6 2.3–3.7 5–10 2.9–5.8 1.5–3.5 5–10 1–2 1–2.2

Using Tukey post hoc comparison test, differences among species indicated by the same superscripted letters were not significant
Chemical properties of good quality meadow hay follow Hejcman et al. (2010, 2012), Hrevusová et al. (Hrevušová et al. 2009) and Tallowin and Jefferson
(1999) and the optimum range for cattle follows Kudrna (1998) and Whitehead (1995)
Calculated by one-way ANOVA, differences among species for all chemical properties were significant (P \ 0.01)

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Table 2 Concentration (mean ± standard error of mean) of NDF neutral detergent fibre, ADF acid detergent fibre, ADL acid detergent lignin
and residual ash in leaf-fodder of the studied species
Species NDF (g kg-1) ADF (g kg-1) ADL (g kg-1) Res. Ash (g kg-1)

Acer platanoides 400 ± 24.3a 313 ± 6.3a,b 102 ± 3.2a,b 37.0 ± 1.9a,b,c
a a,d a
Carpinus betulus 437 ± 16.4 273 ± 10.1 85 ± 6.3 30.1 ± 3.2a,b
a b,c b,c,d
Corylus avellana 476 ± 23.5 342 ± 13.4 163 ± 26.2 43.6 ± 2.7b,c
b e e
Fagus sylvatica 663 ± 11.6 480 ± 7.0 236 ± 9.4 30.0 ± 1.6a,b
a,b c d,e
Fraxinus excelsior 512 ± 77.8 401 ± 13.9 207 ± 14.3 47.7 ± 2.2c
a c c,d,e
Populus tremula 452 ± 27.5 374 ± 24.9 195 ± 22.8 41.1 ± 4.7b,c
Quercus robur 450 ± 10.9a 306 ± 5.3a,b 135 ± 9.6a,b,c 24.8 ± 2.8a
Tilia cordata 409 ± 21.4a 305 ± 7.0a,b 116 ± 4.6a,b 37.3 ± 3.7a,b,c
Ulmus glabra 397 ± 41.2a 243 ± 12.0d 99 ± 13.4a,b 51.8 ± 4.6c
Meadow hay 500–680 340 40
Optimum range 330–450 190–300 Max. 80
Using Tukey post hoc comparison test, differences among species with the same superscripted letter were not significant
Chemical properties of good quality meadow hay follow Worrel et al. (Worrell et al. 1986) and Isselstein et al. (2007) and the optimum range for
cattle follows Kudrna (1998) and Whitehead (1995)
Calculated by one-way ANOVA, differences among species for all chemical properties were significant (P \ 0.01)

into groups with a low concentration of macro-elements
and a high N:P ratio on the right hand side of the diagram
(Fagus, Quercus and Carpinus), with an intermediate
concentration of macro-elements in the middle of the dia-
gram (Corylus, Populus) and with a high concentration of
macro-elements and a low N:P ratio on the left hand side of
the diagram (Ulmus, Fraxinus, Tilia, Acer). Species with a
low variability of leaf chemical properties among the
localities were Acer, Fagus and Quercus as for all four
localities these species plotted close together in the dia-
gram. Species plotting as similar in three localities while
differing in the fourth were Carpinus and Corylus and
finally species with high variability between all the local-
ities were Fraxinus, Populus, Tilia and Ulmus.
Discussion
Forage quality of studied species
The main message of this paper is that there are large dif-
ferences in forage quality among the main broad-leaved
woody species in Central Europe. The species with leaves
of low forage quality are Carpinus, Fagus and Quercus,
species with leaves of intermediate forage quality are
Corylus and Populus, and species with leaves of high forage
quality are Ulmus, Fraxinus, Tilia and Acer. Concentrations
of N in the leaves of all species are in the optimum range for
cattle nutrition and comparable with meadow hay. Nitrogen
requirements of cattle can thus be fully covered using the
leaf hay of all the studied species. Concentrations of P in
leaves of Carpinus and Fagus, in contrast to other species,
are below the optimum range for cattle. Insufficient P
concentration in leaf-fodder represents the real problem for
performance of livestock, especially for lactating cows and
young, quickly growing stock. Although the range of opti-
mum P concentrations in forage for cattle is relatively wide,
lactating cows and young stock require high P concentra-
tions in the upper part of this range, at least above 3 g P
kg-1 (Liebisch et al. 2013). Such high P requirements could
be covered only by the leaf-fodder of Fraxinus and Tilia
collected early in the spring. Relatively high N concentra-
tions in the leaves of all species and also relatively high P
concentrations in several species were given by collection
of leaves shortly after their full flush, as N and P concen-
trations are high in young spring leaves and then decrease
during the vegetation season as they senesce (Kobe et al.
2005). Low P concentrations in the leaves of Carpinus and
Fagus, and therefore their low forage quality, was well
reflected by the N:P ratio which was above 15. In grassland
hay of good forage quality the N:P ratio is generally below
10 (Hejcman et al. 2012).
With the exception of Fagus, concentrations of K in
leaves of the other species were too high for optimum cattle
nutrition. K concentrations too high for livestock nutrition
are also common in the forage from almost all meadow or
crop species, and animals are well adapted to excrete
accessible K through their urine (Kayser and Isselstein
2005). Accessible concentration of K in the forage, if not
above 25 g kg-1, does not represent the main problem in
the use of leaf-fodder for cattle although the optimum
concentration is up to 10 g kg-1.
Concentrations of Ca were excessive in leaf-fodder from
all the species and this may cause relatively severe health

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Fig. 1 Ordination diagram showing results of PCA analysis of
relationships among woody species, N, P, K, Ca and Mg concentra-
tions; N:P, Ca:P and K:(Ca?Mg) ratios, NDF neutral detergent fiber,
ADF acid detergent fiber and ADL acido-detergent lignin. Numbers
indicate the locality: 1 Kozly, 2 Měšice, 3 Běstvina and 4 Mšec
problems connected with decrease in cattle performance if
combined with insufficient P supply. Although both ele-
ments are required by animals in large quantities for bone
growth particularly, their supply must be balanced as
accessible Ca supply decreases absorption of P, thus
causing P deficiency. Excessive Ca concentrations are the
main reason why the Ca:P ratio in the leaf-fodder of all the
species was from two to three times higher than the opti-
mum range for cattle nutrition. Taking into account the
Ca:P ratio, the leaf-fodder from the studied species was of
substantially lower quality than meadow hay with its
optimum ratio.
Concentrations of Mg in leaf-fodder were, with the
exception of slightly higher concentrations in Corylus and
Fraxinus, in the optimum range for cattle nutrition simi-
larly as in the case of meadow hay. Leaf-fodder can thus
fulfill the Mg requirements of cattle.
High K concentration associated with low concentrations
of Ca and Mg resulting in a K/(Ca?Mg) ratio higher than
2.2 (Butler 1963), together with excessive N in forage may
induce grass tetany syndrome (Swerczek 2007). This is not
however the case with leaf-hay forage as the K/(Ca?Mg)
ratio was below the critical value of 2.2 in all species.
The content of NDF was within or below the range cor-
responding to meadow hay from semi-natural grasslands, or
even lower. An appropriate content of NDF stimulates rumen
activity. The important finding is that leaves of woody spe-
cies have a very high content of fibre constituted by the lignin
611
fraction that represents an anti-nutritive, indigestible com-
ponent in the forage. High amounts of leaf-fodder may
restrain forage intake and inhibit digestive enzymes, and
consequently induce adverse effects on animal performance.
From this point of view, leaf-fodder probably only played a
maintenance role enabling cattle to survive the winter but did
not enable live-weight gain. Such fodder during the winter
may also shift calving and lambing to a more favourable
period in comparison to that of contemporary free ranging
cattle and sheep (Balasse et al. 2012). In addition, leaf-fodder
was probably used particularly as a supplement for cattle
during the winter in addition to pasture forage, or as a by-
product of timber harvesting for house construction, the
latter consisting mostly in the use of fir, Abies alba (Akeret
and Rentzel 2001). As we recorded in the Altai Mts., cattle
can also graze senescent grassland biomass during the winter
and can scrape out the biomass hidden under a 20 cm deep
snow layer. Up to the Hallstatt period when the first iron
scythes appeared in Central Europe and thus enabled hay-
making (Hejcman et al. 2013), leaf-fodder probably served
as the main winter food for cattle especially in areas with
deep snow layers. In areas with a snow layer less than 20 cm,
leaf-fodder was probably a supplement to the winter grazing,
or animals could graze on it on their own in the surroundings
of the settlement (Akeret and Jacomet 1997).
Leaf-fodder was probably more widely used for sheep
and goats as they feed naturally more on leaves of shrubs
and trees than cattle and are physiologically adapted to this
(Papachristou et al. 2005). In particular goats may have
mechanisms to attenuate the undesirable effects of lignin
and secondary metabolites (Howe et al. 1988) and their
natural food can contain more than 50 % of leaves of
woody species (Papachristou and Nastis 1993). On the
other hand, the use of leaf-fodder has been widely recorded
for sheep and goats in archaeological localities. This is
because their dung is compact and easily recognisable
(Akeret and Jacomet 1997; Akeret et al. 1999), compared
with cattle dung which forms pats of typical form, but are
usually mixed with the rest of the sediment (Akeret and
Rentzel 2001; Shahack-Gross 2011).
Consequences of leaf-fodder quality for development
of forest vegetation in Holocene
In many European regions, farmers preferred, if possible, to
use Acer, Fraxinus, Ulmus and Tilia for leaf-fodder har-
vesting (Dreslerová 2012). We can conclude that this
preference closely followed their better forage quality in
comparison with other common broad-leaved woody spe-
cies in Central Europe. Similar results, i.e. the substantially
better forage quality of Fraxinus ornus than Carpinus ori-
entalis, were also recorded in Mediterranean (Papachristou
1997). Nowadays, forage quality is analyzed by modern
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analytical methods, but farmers were surely able to recog-
nize the forage quality of different woody species without
any analytical methods, but according to milk yield or live
weight gain of their livestock (our personal experience). For
example the use of Ulmus leaves as supplementary fodder
for cattle in the time of forage shortage was recommended,
for the first time, by Marcus Cato (234–149 B.C.) in his book
De Agricultura (Hooper and Ash 1935). In Iceland in the
6–8th centuries A.D., Ulmus was highly valued as fodder for
cows (Kelly 2000). Farmers were undoubtedly also aware
of other qualities of tree fodder, in particular their medicinal
properties which may, for instance, help to control parasite
infestation of animals (Austad 1988; Musonda et al. 2009;
Martin et al. 2011).
During grazing, livestock make up their diet with spe-
cies of higher forage quality to fulfill their nutritional
requirements. As lactating cows and calves require a high P
supply, they frequently prefer to graze P rich biomass.
Cattle were the most important animals from the first
farmers up to modern times not only in the Czech Republic
(Schibler and Chaix 1995; Kyselý 2012). Phosphorus-poor
leaves of Carpinus and Fagus were thus probably avoided
by cattle during grazing in forests if other woody species
with higher P concentrations and lower lignin content in
leaves were available. As the spread of both species has
occurred since the Bronze Age in Bohemia, the western
part of the Czech Republic (Pokorný and Kuneš 2005;
Chytrý 2012), it is highly probable that this was at least
partly connected to their avoidance by livestock in forests
and also probably by farmers during leaf-fodder harvesting.
Selective browsing and harvesting of Acer, Fraxinus, Tilia
and Ulmus thus probably augmented their decline in the
Bronze and Iron ages and supported the expansion of
Carpinus and Fagus.
Conclusion
Our results suggest that our ancestors’ practice of tree
species exploitation for leaf-fodder in the past was not only
governed by their availability, but also followed the forage
quality of harvested woody species within the actual veg-
etation context. The forage quality of investigated woody
species does not however reach the level of dietary
requirements of contemporary dairy cattle for their satis-
factory performance. Leaf-fodder was thus probably used
as a supplement to biomass grazed during winter and
enabled winter survival without substantial loss of the live
weight of cattle. Species providing the best forage quality
of leaf-fodder were Acer, Fraxinus, Tilia and Ulmus, spe-
cies providing intermediate quality were Corylus and
Populus, and finally species providing very low quality
were Quercus, Carpinus and Fagus.
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Veget Hist Archaeobot (2014) 23:607–613
Acknowledgments The study was funded by a Grant from the
Czech University of Life Sciences Prague CIGA 20114205.
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